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94 Hemolymph Proteins and Functional Peptides: Recent Advances in Insects and Other Arthropods Vol. 1, 2012, 94-110<br />

Muhammad Tufail and Makio Takeda (Eds)<br />

All rights reserved-© 2012 <strong>Bentham</strong> <strong>Science</strong> Publishers<br />

CHAPTER 6<br />

Structures and Functions of Insect Midgut: The Regulatory Mechanisms<br />

by Peptides, Proteins and Related Compounds<br />

Makio Takeda *<br />

Graduate School of Agricultural <strong>Science</strong>, Kobe University, 1-1Rokkodai-cho, Nada-ku, Kobe, 657-8501,<br />

Japan<br />

Abstract: The midgut is an important organ for insects not only because it occupies a large space in their<br />

hemocoel, or because it is a major part of the digestive tract; it also plays critical roles in other physiological<br />

regulation such as metabolism, immune response, homeostasis of electrolytes, osmotic pressure, circulation<br />

and more. Therefore disturbance of these functions could provide a target and strategy for future pest<br />

management practice. Entry of entomopathogenic microorganisms and penetration of pesticides are through<br />

this route, and Plasmodium penetrating through the peritrophic membrane of Anopheline mosquitoes is a<br />

crucial issue for malaria control. The midgut is a complex organ that undergoes a gross transformation at<br />

metamorphosis to support different feeding habits at different developmental stages. It consists of<br />

epithelium and basal lamina with endodermal and mesodermal origins, respectively and the latter is<br />

innervated, i.e., an ectodermal element. We have not yet reached complete elucidation and understanding of<br />

the mechanism of embryonic formation of the organ, the metamorphosis and the regulatory mechanisms for<br />

a variety of midgut house-hold physiological functions. Another complexity comes from the unusual extent<br />

of variabilities that occur in general morphology, egg size/yolk content, feeding habit, mode of<br />

metamorphosis, or kinds of stresses or symbiotic associations that may occur in various orders of insects.<br />

We do not know whether the Drosophila model that provides important knowledge concerning embryonic<br />

formation is applicable to other insects. This review focuses on functions of developmental genes, endocrine<br />

agents, and local secretagogues, that control regulatory mechanisms, particularly peptides secreted from<br />

epithelial paraneurons as well as growth factors and differentiation factors synthesized by non-paraneuronal<br />

midgut cells such as columnar cells, goblet cells, perhaps trachea, hemocytes or muscles or extramidgut<br />

tissues such as the fat body. Stress conditions and metamorphosis, epithelial-mesenchymal interactions and<br />

behavior of adhesion molecules should be included here but currently this aspect has received little<br />

attention.<br />

Keywords: Brain-gut peptides, paraneurons, growth factors, differentiation factors, metamorphosis, gut<br />

contraction, digestion, metabolism, 20-hydroxyecdysone, bombyxin, serotonin, apoptosis, stem cell.<br />

INTRODUCTION<br />

Hormones are circulating agents produced by endocrine glands that control physiological functions in sites<br />

remote from the site of production. The nervous system also produces chemical signals with structures<br />

similar to hormones and secretes them not only to synaptic junctions but also to the general circulation. For<br />

example, adrenalin is a hormone but is also released into synaptic junctions. Ernst and Bertha Scharrer [1]<br />

unified endocrinological and neurological traditions by the integrating concept of “neurosecretion”.<br />

Neurons secrete conventional neurotransmitters (synaptocrinia), neuropeptides (synamptocrinia, paracrinia<br />

and endocrinia) or autocrine agents such as nerve growth factor (autocrinia). Confusions, however still<br />

remain; some non-neural tissues can produce “neuropeptides”. This was originally found in mammalian<br />

intestines, thymus, placenta etc., and the same conditions are found in insects. Some like Pearse [2]<br />

assumed that these “endocrine cells” must have an ectodermal origin and have migrated from the neural<br />

crest to the epithelium, the APUD hypothesis (Amine Precursor-Uptake- Decarboxylation). However,<br />

currently the midgut “endocrine cells,” at least in insects, are considered as having their origin different<br />

from neural tissue [3, 4].<br />

*Address correspondence to Makio Takeda: Graduate School of Agricultural <strong>Science</strong>, Kobe University, 1-1Rokkodai-cho, Nadaku,<br />

Kobe, 657-8501, Japan; Ph/Fax; +81-78-803-5870; Email: mtakeda@kobe-u.ac.jp

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